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Atlantic Geology

Isocrinid crinoids from the late Cenozoic of Jamaica Stephen K. Donovan

Volume 30, Number 3, November 1994 Article abstract Eight species of isocrinines have been documented from the Lower Cretaceous URI: https://id.erudit.org/iderudit/ageo30_3rep02 to Pleistocene of Jamaica. New finds include a second specimen of a Miocene species from central north Jamaica, previously regarded as Diplocrinus sp. but See table of contents reclassified as Teliocrmus? sp. herein. Extant Teliocrinus is limited to the Indian Ocean, although Miocene specimens have been recorded from Japan, indicating a wider distribution during the Neogene. One locality in the early Publisher(s) Pleistocene Manchioncal Formation of eastern Jamaica has yielded three species of isocrinine, Cenocrinus asterius (Linnl), Diplocrinus maclearanus Atlantic Geoscience Society (Thomson) and Neocrinus decorus Thomson. These occur in association with the bourgueticrinine Democrinus sp. or Monachocrinus sp. These taxa are all ISSN extant and suggest a minimum depositional depth for the Manchioneal Formation at this locality of about 180 m. This early Pleistocene fauna 0843-5561 (print) represents the most diverse assemblage of fossil crinoids docu mented from 1718-7885 (digital) the Antillean region.

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Cite this article Donovan, S. K. (1994). Isocrinid crinoids from the late Cenozoic of Jamaica. Atlantic Geology, 30(3), 195–203.

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This article is disseminated and preserved by Érudit. Érudit is a non-profit inter-university consortium of the Université de Montréal, Université Laval, and the Université du Québec à Montréal. Its mission is to promote and disseminate research. https://www.erudit.org/en/ A tlantic G eology 195

Isocrinid crinoids from the late Cenozoic of Jamaica

Stephen K. Donovan Department of Geology, University of the West Indies, Mona, Kingston 7, Jamaica

Date Received April 8, 1994 Date A ccepted A ugust 26, 1994

Eight species of isocrinines have been documented from the Lower Cretaceous to Pleistocene of Jamaica. New finds include a second specimen of a Miocene species from central north Jamaica, previously regarded as Diplocrinus sp. but reclassified as Teliocrinus? sp. herein. Extant Teliocrinus is limited to the Indian Ocean, although Miocene specimens have been recorded from Japan, indicating a wider distribution during the Neogene. One locality in the early Pleistocene Manchioneal Formation of eastern Jamaica has yielded three species of isocrinine, Cenocrirtus asterius (Linne), Diplocrinus maclearanus (Thomson) and Neocrinus decorus Thomson. These occur in association with the bourgueticrinine Democrinus sp. or Monachocrinus sp. These taxa are all extant and suggest a minimum depositional depth for the Manchioneal Formation at this locality of about 180 m. This early Pleistocene fauna represents the most diverse assemblage of fossil crinoids docu­ mented from the Antillean region.

Huit especes d’isocrinines de la periode du Cretace inferieur au Pleistocene de la Jamai'que ont ete documentees. Les nouvelles decouvertes comprennent un deuxieme specimen d’une espece du Miocene du nord central de la Jamai'que, auparavant consideree comme l’espece Diplocrinus, mais reclassifiee en tant que Teliocrinus? aux presentes. Le Teliocrinus existant est limite a l’ocean Indien, meme si on a releve des specimens du Miocene au Japon, ce qui est revelateur d’une distribution plus repandue au cours du Neogene. Un emplacement de la formation de Manchioneal du Pleistocene inferieur dans l’est de la Jamai'que a livre trois especes d’isocrinines : le Cenocrinus asterius (Linne), le Diplocrinus maclearanus (Thomson) et le Neocrinus decorus (Thomson). Ceux-ci sont presents en association avec les bourgueticrinines Democrinus ou Monachocrinus. Ces taxons sont tous existants et ils permettent de supposer que la formation de Manchioneal aurait une profondeur minimale de sedimentation d’environ 180 m a cet endroit. Cette faune du Pleistocene inferieur represente l’assemblage le plus diversify de crinoi'des fossiles documentes dans la region des Antilles. [Traduit par la redaction]

Introduction S ystematic P aleontology

Few fossil crinoids are known from Jamaica; nevertheless, Class Crinoidea J.S. Miller, 1821 they are probably the best known fauna of this group from the Subclass Articulata von Zittel, 1879 Antillean islands. Most of the described taxa are based on just Order Isocrinida Sieverts-Doreck, 1952 one or a few specimens. Of the eight species hitherto recog­ Suborder Isocrinina Gislen, 1924 nized (Table 1), five are isocrinines; all of these are recorded Family Isocrinidae Gislen, 1924 from a single columnal or pluricolumnal only and each from a different locality and horizon. It is therefore considered signifi­ Genus Neocrinus Thomson, 1864 cant to report further specimens from two of these horizons, Neocrinus decorus Thomson, 1864 generally showing superior preservation to those originally de­ Figure 2.1,2.2,2.4,2.6,2.7,2.9-2.11 scribed. The discovery of a moderately rich crinoid locality in the early Pleistocene of eastern Jamaica, including three spe­ Material: Twelve columnals and pluricolumnals, BMNH EE cies of isocrinine and a bourgueticrinine, represents the first 5052, EE 5053, EE 5057[2], EE 5060, EE 5062 (Fig. 2.7,2.10, site on the island to yield more than one species of crinoid and 2.11), EE 5064 (Fig. 2.1, 2.4), EE 5065[2] (Fig. 2.2, 2.6), EE has produced more specimens than all of the other Cenozoic 5065[3] (Fig. 2.9), EE 5067-EE 5070. sites combined. Terminology of the crinoid stem used herein follows Moore Occurrence: Cliffs and associated fallen blocks exposed near etal. (1968), Roux (1977), Ubaghs (1978) and Webster (1974). the mouth of the Christmas River, parish of Portland, eastern The classification of articulate crinoids follows Simms (1988). Jamaica (Fig. 1.1; GR 798 421, old 1:50,000 series, sheet M, The protocol of open nomenclature follows Bengtson (1988). ‘Port Antonio’). Upper Coastal Group, Manchioneal Forma­ The specimens described herein are deposited in the Natural tion. Early Pleistocene. History Museum, London (BMNH) and the Florida Museum of Natural History (UF).

Atlantic Geology 30,195-203 (1994) 0843-5561/94/030195-12S2.35/0 196 D onovan

Table 1. Fossil crinoids of Jamaica (after Donovan, 1989; Donovan etal., 1993, 1994; Dixon etal., 1994). * = discussed herein.

Order ISOCRINIDA Suborder COMATULIDINA Incertae familiae Brachial or pinnular ossicle late Oligocene Brachial ossicles indet. late Cretaceous Suborder ISOCRININA Family ISOCRINIDAE * Neocrinus decorus Thomson early Pleistocene Family CAINOCRINIDAE * Teliocrinus? sp. early Miocene Family ISSELICRINIDAE Austinocrinus sp. early Aptian cf. Cenocrinus asterius (Linne) early Pleistocene * Cenocrinus asterius (Linne) early Pleistocene * Diplocrinus maclearanus (Thomson) early Pleistocene Isselicrinus cubensisl (Valette) early Eocene Incertae familiae Isocrinid sp. indet. middle Eocene Suborder BOURGUETICRININA Family BATHYCRINIDAE * Democrinus sp. or Monachocrinus sp. early Pleistocene Order ROVEACRINIDA Family SACCOCOMIDAE Applinocrinus cretacea (Bather) Maastrichtian

Description: For a description of the stem of extant N. decorus, Formation (Pliocene) is apparent at Christmas River, indicat­ see Donovan (1984). ing that the crinoid-rich beds are low in the Manchioneal For­ mation. Many of the specimens from Christmas River have been Remarks: The Pleistocene columnals illustrated herein com­ collected from fallen blocks. pare closely with those of extant N. decorus (for example, com- Evidence that pluricolumnals may have lain on the seaf­ pareFig. 2.1,2.4 herein with Roux, 1977,figs. 7C, 19;Donovan, loor for some time between death and final burial is given by an 1984, pi. 74, fig. 2). Even though the trabeculae are encrusted epifaunal, serpulid worm tube (Fig. 2.11). This encrusts two by microspar, the rectilinear stereom of the areola petals is iden­ adjacent columnals, which presumably would have been im­ tical to that of living examples (compare Fig. 2.6 with Roux, practicable when the crinoid was alive due to the flexibility of 1977, fig. 5A; Donovan, 1984, pi. 75, fig. 7). The symplectial the column (compare with Donovan, 1993, fig. 2C). Although facet of N. decorus is easily discerned from those of Diplocrinus massive encrusting epibionts are not uncommon on Paleozoic maclearanus and Cenocrinus decorus (see below) by its dis­ crinoids (Meyer and Ausich, 1983), presumably immobilzing tinctive, teardrop-shaped areola petals. Extant N. decorus oc­ the column in the region of attachment, the Pleistocene speci­ curs in the tropical western Atlantic, including off Jamaica men is small and fragile. It thus suggests that the column was (Meyer etal., 1978, p. 425). not capable of flexure (=dead) when encrusted. The cliffs in the Manchioneal Formation near the mouth of the Christmas River are only the second moderately rich, Family Cainocrinidae Simms, 1988 crinoid-bearing succession to be identified from the Jamaican rock record, the fauna consisting of Cenocrinus asterius (Linne), Genus Teliocrinus Doderlein, 1912 Diplocrinus maclearanus (Thomson), Neocrinus decorus and Teliocrinus? sp. Democrinus sp. or Monachocrinus sp. (see below). The first Figure 3 such locality, at San San Bay, parish of Portland (Robinson, 1993 Diplocrinus sp., Donovan etal., pp. 126-127, fig. 4. 1969), is in the same unit, but is unfortunately now concealed by a retaining wall (Donovan, 1989). Columnals were quite com­ Material: Two internodal columnals, UF 67000 (Fig. 3) and mon at San San Bay (Robinson, 1969,1971), but the taxonomic UF 38939 (Donovan etal., 1993, fig. 4). diversity of this locality is unknown. The present author and co-workers have examined the Manchioneal Formation for bra- Occurrence: Disused quarry about 5 km west of Duncans po­ chiopods at a number of localities, but have not previously noted lice station, on the southern side of the main A1 (north coast) associated crinoids. The contact with the underlying Bowden road, parish of Trelawny, central north Jamaica (Fig. 1.2; ap- A tlantic G eology 197

Fig. 1. Locality maps. The inset map of Jamaica shows the positions of main maps (1) and (2). (1) Christmas River area, eastern Jamaica (rivers other than the Christmas River are not shown for simplicity). (2) Duncans-Falmouth area, central north Jamaica. Key: filled circles = crinoid localities, stippled line = coast, thick lines = main roads, thin line marked cr = Christmas River. proximate GR 349 570, old 1:50,000 series, sheet C, ‘Montego lumen diameter = 0.35 mm; UF 38939 - KH = 1.6 mm, KD = Bay-Falmouth’). SignMember(7), Montpelier Formation, White 5.7 mm. Limestone Group. Miocene, probably early Miocene (E. Robinson, personal communication). Remarks: This Miocene species was originally described on the basis of UF 38939 (Donovan et al., 1993), a broken intem- Description: (Revised after Donovan et a l, 1993). Columnal odal. UF 67000 is better preserved (Fig. 3), being unbroken, outline weakly pentastellate with strongly rounded angles. Axial with the lumen exposed and at least some petals reasonably canal small, central and circular in outline. Articulation clean (Fig. 3.3). The areola petals are now recognized to be symplectial, arranged about five slender, elongate, lensoid are­ open, not closed as was originally reported; the crenulae of ad­ ola petals that correspond to the columnal angles. Areola petals jacent petals are apparently separate (Fig. 3.1, 3.3) (Donovan separate, except adjacent to lumen, and open. Crenulae perpen­ etal., 1993). dicular to subperpendicular to circumference of areola petals. The Miocene columnals compare reasonably well with those Crenulae short, unbranched, 6 to 7 culmina along each side of of extant species of Diplocrinus, such as D. maclearanus, D. the areola petals, with crenulae of adjacent petals separate. Tri­ altemicirrus and D. wyvillethomsoni (Roux, 1977; Macurda and angular naked zones occur adjacent to the circumference in Roux, 1981), conspicuous similarities include the slender, elon­ interpetaloid zones. Latus gently convex and unsculptured. gate, open petals. The triangular naked zone at the circumfer­ ence in the interpetaloid zones are similar to those in D. Dimensions: UF 67000 - columnal height (KH) = ca. 1.5 mm, altemicirrus (Roux, 1977, figs. 7F, 16) and D. maclearanus columnal diameter (KD) = 6 mm, facet diameter = 5.8 mm, (Macurda and Roux, 1981, pi. 15.4-15.7). However, all three NVAONOQ 8 6 1 A tlantic G eology 199

Fig. 3. Teliocrinus! sp. from the Miocene of Jamaica, UF 67000. (1) Articular facet, x9. (2) Lateral view, x9. (3) Detail of an areola petal, x l8. Scanning electron micrographs of specimens coated with 60% gold-palladium.

Recent species differ from the fossil in that the culmina are the Pacific during the Miocene and became an exposed land relatively narrower (and are thus more plentiful) in the latter, mass in the Plio-Pleistocene (Pindell and Barrett, 1990, p. 419). whereas culmina of adjacent petals remain separate, that is, there is an interpetaloid groove. The three extant species mentioned Family Issclicrinidae Klikushin, 1977 above have only 4 to 5 culmina along each side of the areola petals, but the Miocene species has 6 to 7 (Fig. 3.1, 3.3; Donovan Genus Cenocrinus Thomson, 1864 et al., 1993, fig. 4). As discussed above, the distinguishing fea­ Cenocrinus asterius (Linne, 1767) tures of Diplocrinus include the lanceolate areola shape, the Figure 4 open crenularium with less than 8 crenulae in each petaloid zone, and an undifferentiated interpetaloid zone without an Material: An internodal, BMNH EE 5050 (Fig. 4). Two interpetaloid groove (Roux, 1977, table 2). The presence of more pluricolumnals, BMNH EE 5051, EE 5055, are also tentatively than 10 culmina per petal and interpetaloid grooves in the Ja­ included in this species. maican specimen suggests that this taxon is closer to Teliocrinus Doderlein, a genus otherwise only known from the Recent of Occurrence: Cliffs and associated fallen blocks exposed near the Indian Ocean (Roux, 1977, table 1; Rasmussen, 1978, p. the mouth of the Christmas River, parish of Portland, eastern T860) and the Miocene of Japan (Oji, 1990). Comparison with Jamaica (Fig. 1.1; GR 798 421, old 1:50,000 series, sheet M, the symplectial articulations of the type species, T. springeri ‘Port Antonio’). Manchioneal Formation, Upper Coastal Group. A.H. Clark, 1909, shows them to be similar (Roux, 1977, figs. Early Pleistocene. 7E, 15). This perhaps suggests that the Jamaican fossil speci­ mens represent a new species of Teliocrinus, although this ge­ Description: Pentalobate internodal columnal with strongly neric identification is not yet certain. It is desirable that more is rounded angles. Lumen central, small and circular. Articula­ known of the skeleton of the Jamaican species before it is for­ tion symplectial, arranged about five moderately broad, lensoid mally named. Dr. T.K. Baumiller (written communication) has areola petals that correspond to the columnal angles. Petals sepa­ also noted a similarity between the Jamaican species and rate, divided centrally by a prominent perilumen. Crenulae Isocrinus sp. from the Miocene of Japan (Oji, 1990, fig. 6.7, mainly perpendicular to circumference of areola petals. Crenulae 6.8). short, unbranched, 6 to 7 culmina along each side of areola Oji (1990) noted that extant Teliocrinus is limited to the petals, with crenulae of adjacent petals coalescing towards the tropical Indian Ocean, but it did extend into the western Pacific center. Triangular naked zones occur adjacent to the circumfer­ during the Miocene. If the present identification is correct, in ence in interpetaloid zones. Columnal low. Latus slightly con­ the Miocene its range also extended into the Caribbean. This vex and unsculptured. was facilitated by the absence of the Central American land bridge, which started to interrupt deep water circulation from

Fig. 2. Early Pleistocene crinoid columnals and pluricolumnals from Jamaica. (1, 2, 4, 6, 7, 9-11) Neocrinus decorus Thomson, 1864. (1, 4) BMNH EE 5064. (1) Articular facet, xl4. (4) Detail of an areola petal, x34. (2, 6) BMNH EE 5065[2], (2) Articular facet, xl4. (6) Stereom trabelculae of areola petal overgrown by microspar, x850. (7, 10, 11) BMNH EE 5062. (7) Cirral facet of nodal, x32. (10) Pluricolumnal, x7.5. (11) Serpulid worm tube encrusting pluricolumnal, x39. (9) BMNH EE 5065[3], distal cryptosymplectial articulation of a broken nodal columnal, xl4. (3, 5 ,8) Diplocrinus maclearanus (Thomson). (3) BMNH EE 5057[1], articular facet, xl2. (5) BMNH EE 5058, pluricolumnal (intemoditaxis), with cryptosymplectial articulation at top, x8. (8) BMNH EE 5059, apparent growth deformation of facet, xl4. Scanning electron micrographs of specimens coated with 60% gold-palladium. 2 0 0 D onovan

mas River, both basal sections of the Manchioneal Formation, suggesting that this unit is a shallowing-upwards sequence (Donovan and Embden, in press).

Genus Diplocrinus Doderlein, 1912 Diplocrinus maclearanus (Thomson, 1877) Figure 2.3,2.5, 2.8

Material: Five columnals and pluricolumnals, BMNH EE 5057[1] (Fig. 2.3), EE 5058 (Fig. 2.5), EE 5059 (Fig. 2.8), EE 5065[1], EE 5066. A further specimen, BMNH EE 5054, is only tentatively included in this species.

Occurrence: Cliffs and associated fallen blocks exposed near the mouth of the Christmas River, parish of Portland, eastern Jamaica (Fig. 1.1; GR 798 421, old 1:50,000 series, sheet M, ‘Port Antonio’). Manchioneal Formation, Upper Coastal Group. Early Pleistocene.

Description: For a description of the stem of extant D. maclearanus, see Macurda and Roux (1981).

Remarks: Roux (1977, table 2) considered the distinguishing features of Diplocrinus, based on D. alternicirrus (Carpenter) 0 2 and D. wyvillethomsoni (Jeffreys), to be the lanceolate shape of I______i______I the areola petals, the open crenularium with less than 8 crenulae mm in each petaloid zone, and an undifferentiated interpetaloid zone without an interpetaloid groove. These features are all apparent Fig. 4. Cenocrinus asterius (Linne, 1767) from the early Pleistocene in the figured articular facet (Fig. 2.3), which agrees well with of Jamaica, articular facet, BMNH EE 5050. Camera lucida drawing. that of D. maclearanus (Macurda and Roux, 1981, pi. 15, figs. 4-7). Meyer et al. (1978, p. 426) did not note the occurrence of Remarks: Donovan (1989) identified cf. Cenocrinus asterius this species in Jamaican waters at the present day, but it is broadly from the Manchioneal Formation at San San Bay (Table 1), distributed in the tropical western Atlantic. based on a single, poorly preserved columnal. Comparison of BMNH EE 5059 is a pluricolumnal with one exposed facet BMNH EE 5050 with illustrations of the articular facet in ex­ that compares with D. maclearanus. However, the other facet tant C. asterius (Roux, 1977, fig. 3E, F, 13; Donovan, 1989, (Fig. 2.8) appears to have a deformity in which two petals are fig. 2C, D, F; Donovan et al., 1993, fig. 3.7) suggests that it is growing particularly close together so that their crenulae are in conspecific, although the specimens illustrated by Roux appear close association. to have slightly more angular areola petals. Extant C. asterius is known to occur at water depths of 183 Incertaefamiliae to 585 m in the western Atlantic, including offshore Jamaica (Meyer et al., 1978, p. 424). This suggests a minimum depth of Isocrinids gen et sp. indet. deposition for the Manchioneal Formation at Christmas River of about 180 m; the other two nominal species from this local­ Material: Six pluricolumnals, BMNH EE 5056, EE 5061, EE ity are known from minimum depths of 154 (N. decorus) and 5071-EE 5074. 187 m (D. maclearanus) (Meyer et al., 1978, pp. 425, 426). Even if these depth minima are inaccurate, extant stalked Occurrence: Cliffs and associated fallen blocks exposed near crinoids rarely extend into water depths of less than 150 m (Lane the mouth of the Christmas River, parish of Portland, eastern and Webster, 1980, p. 145). This depth range is in agreement Jamaica (Fig. 1.1; GR 798 421, old 1:50,000 series, sheet M, with estimates based on sedimentology and brachiopods (E.N. ‘Port Antonio’). Upper Coastal Group, Manchioneal Forma­ Doyle, personal communication), echinoids (Donovan and Em- tion. Early Pleistocene. bden, in press) and ‘deep-water alcyonarian corals’ (Robinson, 1969, p. 9). However, it is at variance with data from ostracodes Remarks: The articular facets on these specimens are poorly (van den Bold, 1971), which suggest that most deposition of preserved, but they probably belong to one of the three species the Manchioneal Formation occurred at less than 100 m water documented from Christmas River (see above). depth. Crinoids have only been found at San San Bay and Christ­ A tlantic G eology 2 0 1

Fig. 5. Democrinus sp. or Monachocrinus sp. from the early Pleistocene of eastern Jamaica. (1-3) BMNH EE 5136. (1) Articular facet, x40. (2) Oblique view of articular facet and latus, x36. (3) Detail of the fulcral ridge, xl25. (4) BMNH EE 5137, lateral view, x21. All scanning electron micrographs of specimens coated with 60% gold-palladium.

Suborder Bourgueticrinina Sicverts-Doreck, 1953 Description: Articular facet lenticular in outline, bifascial, with Family Bathycrinidac Bather, 1899 synarthrial ridge corresponding to long axis. Synarthrial ridge with axial groove flanked by offset knobs separated by large Democrinus sp. or Monachocrinus sp. pores. Facet flanking synarthry relatively depressed. Center of Figure 5 columnal strongly depressed, shaped like broad figure ‘8’, with rounded(?) axial canal at base of depression. Stereom of facet Material: Two columnals, BMNH EE 5136, EE 5137. EE 5136 galleried(?), knobs of synarthrial ridge imperforate. exposes a well-preserved articular facet (Fig. 5.1, 5.3), although Columnal higher than wide, with unsculptured, planar the latus is largely obscured (Fig. 5.2). EE 5137 has poorly pre­ latera that are slightly convex adjacent to latera. Long axes of served articula, but a well-exposed latus (Fig. 5.4). latera offset within any columnal.

Occurrence: From fallen blocks exposed near the mouth of the Remarks: These columnals are typical of bathycrinids, both in Christmas River, parish of Portland, eastern Jamaica (Fig. 1.1; their facet and latus geometry (for example, compare with il­ GR 798 421, old 1:50,000 topographic series, sheet M, ‘Port lustrations in Macurda and Meyer, 1975, pi. 2, figs. 4-7; 1976, Antonio’). Manchioneal Formation, Upper Coastal Group. Early pi. 1). Their occurrence in the early Pleistocene of the Carib­ Pleistocene. bean is not unexpected, as the only extant bourgueticrinines in 2 0 2 D onovan the region belong to this family (Meyer et al., 1978, pp. ophiuroid ossicles from the Oligocene of Jamaica. Caribbean Jour­ 426-427). Unfortunately, without the evidence of the crown, it nal of Science, 30, pp. 143-145. is impossible to classify these columnals with confidence below D o derlein , L. 1912. Die gestielten Crinoiden der deutschen Tiefsee-Expedition. Wissenschaftliche Ergebnisse Deutsche familial level. However, the extant Caribbean taxa belong to Tiefsee-Expedition 1898-1899,17, pp. 1-34. Gustav Fischer, Jena. just two genera, Democrinus Perrier and Monachocrinus A.H. D onovan, S.K. 1984. Stem morphology of the Recent crinoid Clark, and it seems probable that these fossil columnals belong Chladocrinus (Neocrinus) decorus. Palaeontology, 27, pp. 825- to one of these taxa. The Caribbean bathycrinids have the fol­ 841. lowing depth ranges (Meyer et al. ,1978): Democrinus rawsonii ------1989. An isocrinid (Echinodermata: Crinoidea) from the Lower (Pourtales), 66 to 652 m; D. conifer (A.H. Clark), 170 to 1750 Pleistocene of Portland, eastern Jamaica. Journal of the Geologi­ m; D. brevis (A.H. Clark), 210 to 878 m; and Monachocrinus cal Society of Jamaica, 25 (for 1988), pp. 3-7. caribbeus (A.H. Clark), possibly 421 to 1857 m. On the basis ------1993. Contractile tissues in the cirri of ancient crinoids: crite­ of these figures, it is tentatively suggested that the Manchioneal ria for recognition. Lethaia, 26, pp. 163-169. D onovan, S.K. and E mbden, B.J. In press. Early Pleistocene echinoids Formation specimen is most likely to represent a Democrinus of the Manchioneal Formation, Jamaica. Journal of Paleontology. sp. D onovan, S.K., G ordon, C.M., V eltkamp, C.J., and Scott, A.D. 1993. The column of bathycrinids tends to be divided into a short Crinoids, asteroids and ophiuroids in the Jamaican fossil record. proxistele, with low columnals articulating synostosially, a long In Biostratigraphy of Jamaica. Edited by R.M. Wright and E. mesistele with relatively few, tall columnals, and a short dististele Robinson. Geological Society of America Memoir, 182, pp. adapted as a radicular attachment structure (Donovan and 125-130. Pawson, research in progress). The mesistele may be further D onovan, S.K., M iller, S.A., G raham, A.P., and D ixon, H.L. 1994. subdivided into more proximal, cylindrical columnals and more New fossil crinoids from Jamaica. Journal of Paleontology, 68, distal columnals with strongly elliptical articular facets, which pp. 842-845. give the columnal a ‘bone-like’ appearance. The present speci­ G islen, T. 1924. Echinoderm studies. Zoologiska Bidrag fr&n Uppsala, 9, 316 p. mens obviously belong to the more distal mesistele. K likushin, V.G. 1977. Morskiye lilii roda Isselicrinus. Paleontologicheskiy Zhumal 11, pp. 87-95. [In Russian; English A cknowledgements translation 1978. Sea lilies of the genus Isselicrinus. Paleonto­ logical Journal 11, pp. 87-95.] I thank Roger W. Ported (Florida Museum of Natural His­Lane, N.G. and W ebster, G.D. 1980. Crinoidea. In Echinoderms: Notes tory) for collecting and loaning the second isocrinid columnal for a Short Course. Edited by T.W. Broadhead and J.A. Waters. from the Montpelier Formation. Cornelis J. Veltkamp (Univer­ University of Tennessee Studies in Geology, 3. Knoxville, Ten­ nessee, pp. 144-157. sity of Liverpool) kindly took the SE micrographs in Figures 2 L inne, C. 1767. Systema Naturae. 12th edition, v. 1, pt. 2. Laurentius and 3. Figure 5 was photographed in the EM Laboratory, Na­ Salvius, Holmiae, pp. 533-1327. [Not seen] tional Museum of Natural History, Smithsonian Institution M acurda, D.B., Jr. and M eyer, D.L. 1975. The microstructure of the (USNM), and I thank Walter R. Brown and Susann G. Braden crinoid endoskeleton. University of Kansas Paleontological Con­ for their assistance. This research was initiated during a period tributions, Paper 74, 22 p. of study leave at the Department of Earth Sciences, University ------1976. The morphology and life habits of the abyssal crinoid of Liverpool, supported by a Study and Travel Grant from the Bathycrinus aldrichianus Wyville Thomson and its paleontologi­ University of the West Indies and an award from the Leverhulme cal implications. Journal of Paleontology, 50, pp. 647-667. Trust, and was completed during the tenure of a Senior Re­ M acurda, D.B., J r. and Roux, M. 1981. The skeletal morphology of the isocrinid crinoids Annacrinus wyvillethomsoni and search Fellowship at the USNM, all of which are gratefully ac­ Diplocrinus maclearanus. Contributions from the Museum of knowledged. I thank William I. Ausich (The Ohio State Uni­ Paleontology, University of Michigan, 25, pp. 169-219. versity), Tomasz K. Baumiller (Harvard University), David L. M eyer, D.L. and A usich, W.I. 1983. Biotic interactions among Recent Meyer (University of Cincinnati) and Michael J. Simms and among fossil crinoids. In Biotic Interactions in Recent and (Cheltenham and Gloucester College of Higher Education, U.K.) Fossil Benthic Communities. Edited by M.J.S. Tevesz and P.L. for their constructive comments on earlier versions of this pa­ McCall. Plenum, New York, pp. 377-427. per. M eyer, D.L., M essing, C.G., and M acurda, D.B., Jr. 1978. Zoogeog­ raphy of tropical western Atlantic Crinoidea (Echinodermata). Bulletin of Marine Science, 28, pp. 412-441. B ather, F.A. 1899. A phylogenetic classification of the Pelmatozoa. M iller, J.S. 1821. A Natural History of the Crinoidea or Lily-shaped Report of the British Association for the Advancement of Science Animals, with Observations on the Genera Asteria, Eurayle, for 1898, pp. 916-923. Comatuia and Marsupites. Bryan and Co., Bristol, 150 p. B engston, P. 1988. Open nomenclature. Palaeontology, 31, pp. 223-227. M oore, R.C., J effords, R.M., and M iller, T.H. 1968. Morphological B old, W.A., V an D en. 1971. 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